Research on the inheritance of seed dormancy in lettuce (Lactuca sativa L.) and selection for non-dormancy

Summary Inheritance of dormancy and the results of selection of non-dormant genotypes in segregating populations of lettuce were investigated. Diallel crosses were therefore carried out between two dormant (DOR) and two non-dormant (NDOR) cultivars. F₁, F₂ and F₃ populations were analysed.Environmental variation for dormancy usually was large. The mean germination time (GT) of F₁ seeds from the NDOR % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaciaacaqabeaadaqaaqaaaKaaafaafaqabeGaba% aajqgaa+FaaiaadIhaaeaacaWG4baaaaaa!3B90!\[\begin{array}{*{20}c}x \\x \\\end{array}\]DOR crosses was often intermediate to the GT of the NDOR and DOR parent. The mean GT of F₁ seeds from DOR % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaciaacaqabeaadaqaaqaaaKaaafaafaqabeGaba% aajqgaa+FaaiaadIhaaeaacaWG4baaaaaa!3B90!\[\begin{array}{*{20}c}x \\x \\\end{array}\]DOR crosses equalled the mean GT of the parents; the same applies for the NDOR % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaciaacaqabeaadaqaaqaaaKaaafaafaqabeGaba% aajqgaa+FaaiaadIhaaeaacaWG4baaaaaa!3B90!\[\begin{array}{*{20}c}x \\x \\\end{array}\]NDOR crosses. No differences between reciprocals were observed and neither were such differences found for F₂ populations. F₂ populations from DOR % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaciaacaqabeaadaqaaqaaaKaaafaafaqabeGaba% aajqgaa+FaaiaadIhaaeaacaWG4baaaaaa!3B90!\[\begin{array}{*{20}c}x \\x \\\end{array}\]DOR crosses showed no significant segregation of rapidly germinating seeds and in F₂ populations from NDOR % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaciaacaqabeaadaqaaqaaaKaaafaafaqabeGaba% aajqgaa+FaaiaadIhaaeaacaWG4baaaaaa!3B90!\[\begin{array}{*{20}c}x \\x \\\end{array}\]DOR crosses no accumulation of genes for long GT occurred. In F₂ populations from NDOR % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaciaacaqabeaadaqaaqaaaKaaafaafaqabeGaba% aajqgaa+FaaiaadIhaaeaacaWG4baaaaaa!3B90!\[\begin{array}{*{20}c}x \\x \\\end{array}\]DOR crosses usually 40–60% of the seeds germinated as rapidly as the seeds of the NDOR parents. Only one gene (D) could be responsible for the difference in dormancy behaviour of the DOR and NDOR cultivars. The behaviour of the F₃ lines from various NDOR % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaciaacaqabeaadaqaaqaaaKaaafaafaqabeGaba% aajqgaa+FaaiaadIhaaeaacaWG4baaaaaa!3B90!\[\begin{array}{*{20}c}x \\x \\\end{array}\]DOR crosses supports this hypothesis. A regression of F₃ means on the value of F₂ seeds for GT of various NDOR % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabaGaciaacaqabeaadaqaaqaaaKaaafaafaqabeGaba% aajqgaa+FaaiaadIhaaeaacaWG4baaaaaa!3B90!\[\begin{array}{*{20}c}x \\x \\\end{array}\]DOR crosses showed that h²-narrow usually was rather high for most crosses implying that selection for non-dormancy can be carried out in F₂ populations.     

Analysis of the differences in growth between five lettuce cultivars marking the development in lettuce breeding for winter production

Summary Since 1950 growth of lettuce for winter production has been improved considerably by breeding. It has been investigated whether this improvement must be attributed to a more rapid leaf production, to larger leaves or to both. To this end the differences in growth between the cultivars Meikoningin. Proeftuin's Blackpool, Rapide, Deci-Minor and Valentine were analysed at a range of temperatures (10, 14, 17 and 20°C) under natural light conditions in autumn. The last four cultivars represent successive improvements in growth of lettuce for winter production.From the experiments it can be concluded that the improvement in growth of lettuce for winter production was initially brought about through increasing the rate of leaf production and there-after through increasing leaf size. At the same time the heat requirement for optimal growth has become higher.     

Genetics of nitrate content in lettuce

Summary Adverse effects on human health makes the high nitrate content frequently found in lettuce (Lactuca sativa L.) grown under low light conditions an undesirable trait. Efforts have been made to breed cultivars with a reduced capacity for nitrate accumulation. In this study components of variance for nitrate content were estimated in F₂ and F₃ generations of ten lettuce crosses. Additive genotypic variances (A) were estimated from F₃ variance components and from the covariance between F₂ plants and corresponding F₃ lines. Estimates of wide sense heritability of the F₂ from crosses between a high nitrate genotype and four low nitrate genotypes ranged from 0.44 to 0.74 and the estimates for A ranged from 0.25 to 0.40 g·l^-1. Estimated wide sense heritabilities of F₂'s from six crosses involving two low nitrate parents ranged from 0.15 to 0.52. The parents of four of the low nitrate crosses showed relatively large effects of genotype x environment (GE) interaction in successive experiments: the nitrate content of the parents reacted differently to environmental changes between experiments. Estimates of A for crosses between low nitrate genotypes without large effects of GE interaction ranged from 0 to 0.19 g·l^-1. The estimated probability of selecting transgressive low nitrate lines in the progeny of a cross between a high and a low nitrate genotype was low (P=0.002–0.039), indicating that large populations should be evaluated to combine the positive traits of modern high nitrate cultivars with low nitrate content from genotypes not adapted to modern cropping practices. In the progenies from crosses between two low nitrate genotypes without important GE effects, only low estimates of the probability of obtaining transgressive low nitrate lines were obtained (P=0.04–0.06). With the growth conditions used in this study, the probability of selecting lines with a nitrate content compatible under all winter conditions with the proposed future maximum permissible level of 2.5 g nitrate per kg fresh matter is low. Therefore the solution of this problem should be found in a combination of low nitrate cultivars and cultural measures that reduce the nitrate content of the crop.     

The inheritance of nitrate content in lettuce (Lactuca sativa L.)

Summary The inheritance of nitrate content in lettuce was analysed using 16 F₂ populations and three F₃ populations. Frequency distributions of nitrate content in F₂ and F₃ populations were unimodal and symmetrical, indicating a quantitative inheritance. Both significant positive and negative deviations of the F₂ mean from the mid-parent value were found, indicating dominance or epistasis. Deviations towards low nitrate content were more frequent than deviations in the other direction. Estimates of heritabilities for nitrate content in the F₂ populations ranged from 18% to 69% and were in most cases above 50%. Crosses between low nitrate cultivars did not have lower estimates of heritability in the F₂ than crosses between cultivars with larger differences in nitrate content. In one case a genotype x experiment interaction for nitrate content of parental cultivars was found.Three F₃ populations of crosses between cultivars with low nitrate content were analysed. Estimates of heritabilities for F₃ line means ranged from 78% to 91% and estimates of the genetic standard deviation of nitrate content in unselected advanced generations ranged from 0.24–0.33 g kg^–1. The estimates of heritabilities and of genetic variation in advanced generations offer good prospects of selection of low nitrate genotypes in lettuce. A comparison of efficiency of selection in the F₂ generation and F₃ line selection is made.     

Increase of genetic variation in some lettuce cultivars by induction of mutations: A technical note

Summary Colchicine treatments of 13 lettuce cultivars for production of polyploid lettuce also resulted in rapid growing diploid mutants.     

Genetics of nitrate content of lettuce, 1: Analysis of generation means

Summary The genetics of nitrate content in butterhead lettuce (Lactuca sativa L.) was studied using the mean values of five parental genotypes and several generations obtained from crosses between them. One high nitrate parental genotype was chosen and four low nitrate ones. A diallel analysis showed additive genetic effects to be the major source of variation in generation means. Estimates of additive genetic effects differed significantly between experiments, indicating genotype x experiment interactions. Effects of dominance were relatively small. The size and direction of dominance varied between experiments. Reciprocal differences were of very limited size and also varied between experiments. The inheritance of nitrate content in lettuce fitted the additive-dominance genetic model.     

Inheritance of insensitivity of lettuce to a surplus of exchangeable manganese in steam-sterilized soils

Summary After steam sterilizing of certain glasshouse soils a surplus of exchangeable manganese (SEM) occurs. Great differences exist between lettuce genotypes for sensitivity to SEM.Research on the inheritance of insensitivity was carried out by intererossing five lettuce genotypes: Lactuca sativa cys. Neptune (sensitive). Plenos. Troppo and Celtuce (all insensitive) and an accession of L. scrriola (insensitive). From analysing various populations (as F₁'s. F₂'s and B₁'s) it appeared that different numbers of genes for insensitivity were present in the parents varying from one gene in Plenos and Troppo, two genes in L. serriola to possibly four genes in Celtuce. Genes in the three first mentioned parents were linked in repulsion phase and a chromosome map for these genes was made.     

Inheritance of partial resistance to Myzus persicae in lettuce

Summary The genetics of partial resistance of lettuce to Myzus persicae was studied using F₁ and F₂ generations of two crosses between a susceptible and partially resistant accession (Norden x Batacer and Liba x Norden) and three crosses in which both parents were partially resistant (Batavia la Brillante x Batacer, Batacer x Liba and CGN4741 x Batacer). Partial resistance to M. persicae inherited quantitatively, without important dominance effects. Only in the cross Batacer x Liba were significant departures of the F₁ and F₂ from the midparent found, which were probably caused by epistatic effects. Reciprocal F₁s had similar resistance levels, indicating the absence of cytoplasmic or other maternal effects. Estimates of broad-sense heritability ranged from 0.34 to 0.61. The results indicated that lines with an improved resistance level can be obtained from crosses between partially resistant accessions, preferably by line selection or the application of indirect marker aided selection.Abbreviations PR partial resistance, partially resistant - S susceptibility, susceptible     

The identification and characteristics of field resistance to lettuce downy mildew (Bremia lactucae Regel)

Summary Eighty-one accessions of three Lactuca species which showed no recognisable race specific resistance to Bremia lactucae when tested in the laboratory as seedlings, exhibited different degrees of susceptibility when exposed to natural field infection. As a group, crisp genotypes had less mildew and a slower rate of disease development than other types of lettuce. Wild forms of Lactuca sativa and Lactuca serriola were particularly susceptible. In a further trial, the low field susceptibility of three lettuce cultivars (Iceberg, Batavia blonde de Paris and Grand Rapids) was confirmed. Disease development on cv. Iceberg was compared to that on the highly susceptible cv. Hilde in experiments where the two cultivars were grown either in close proximity or in isolation. The absolute level of attack on cv. Iceberg depended upon disease pressure and differences between the two types only became apparent approximately 8 wk after sowing.     

Sources and inheritance of resistance to stemphylium leaf spot of lettuce

Summary None of the tested cultivars of lettuce was found resistant to Stemphylium leaf spot, a common disease in Israel. Within a Lactuca saligna population collected in wild lettuce in Israel, resistance was traced. Interspecific crosses of L. saligna x L. sativa were made and the mode of inheritance of resistance to this disease was studied. Resistance is apparently controlled by two genes: one dominant (Sm₁) and one recessive (sm₂).Contribution No. 1176-E 1984 series, from the ARO.     

The role of resistance breeding in the intergrated control of downy mildew (Bremia lactucae) in protected lettuce

Summary Over the last 30 years, six resistance alleles (Dm2, Dm3, Dm6, Dm7, Dm11 andDm16) located in two linkage groups, have contributed to the control of downy mildew in lettuce crops grown under protection (glass or polythene) in northern Europe. More recently, an as yet genetically uncharacterised resistance factor, R18, has also begun to assume importance. The occurrence of the various combinations of these resistance alleles that exist in commercial cultivars has been dictated by the pathotypes ofBremia lactucae used in their selection but also restricted by linkage in repulsion. In the UK, a pathotype ofB. lactucae insensitive to phenylamide fungicides, such as metalaxyl, emerged in 1978 and became prevalent throughout lettuce production areas in subsequent years. The specific virulence of this pathotype was identical to the previously described phenylamide sensitive pathotype NL10 and cultivars carryingDm11, Dm16 or R18 were resistant. Consequently, an integrated control strategy based on the utilisation of metalaxyl on cultivars carryingDm11 provided effective control in UK until 1987 when a new phenylamide insensitive pathotype began to cause problems. The specific virulence of this second pathotype, which was first reported in the Netherlands and France, was identical to the previously described phenylamide sensitive pathotype NL15. Cultivars carryingDm6, Dm16 or R18, but notDm11, were resistant to NL15; consequently an appropriate change in the cultivar recommendations for use in the integrated control strategy was successfully promulgated. It is predicted that variations of this integrated control strategy involving the use of appropriately selectedDm gene combinations may prove effective for some time. This prediction is based on studies of the status of the avirulence loci in the two phenylamide insensitive pathotypes and of the specific virulence characteristics of phenylamide sensitive components of the pathogen population.     

Comparison of sources of resistance to leaf aphids in lettuce (Lactuca sativa L.)

Summary Three lines of lettuce with resistance to Nasonovia ribisnigri, based on the dominant Nr-gene, and four lines selected for partial resistance to Myzus persicae were tested against three species of leaf aphid: N. ribisnigri, M. persicae and Macrosiphum euphorbiae. The effect of the Nr-gene was also studied in a segregating F2 population.In the material tested, resistance to N. ribisnigri was exclusively based on the Nr-gene, lines selected only for resistance to M. persicae showed no resistance to N. ribisnigri. The Nr-gene also induces partial resistance to M. persicae, but the level of this resistance is influenced by other genes, because the lines with Nr-gene differed significantly from each other for reproduction of M. persicae. The Nr-gene had no effect on the resistance of lettuce to M. euphorbiae.In lines with the Nr-gene, levels of resistance to M. persicae and to M. euphorbiae were correlated, suggesting that the resistance may be determined by the same genes. The Nr-line with highest resistance to M. persicae was comparable for this characteristic to the lines selected for resistance to M. persicae.The cultivars Taiwan and Ravel possess a resistance factor to M. euphorbiae that has no effect on M. persicae or N. ribisnigri. Lines selected for resistance to M. persicae also showed partial resistance to M. euphorbiae. Based on the present results no conclusions can be drawn whether this resistance is based on the same genes that provide resistance to M. persicae, or on a resistance factor comparable to that found in Taiwan and Ravel.     

Interactions between plant and aphid genotypes in resistance of lettuce to Myzus persicae and Macrosiphum euphorbiae

Summary Six lettuce lines, representing two types of resistance to the green peach aphid, Myzus persicae, and a control line with high susceptibility to M. persicae were tested for resistance to six different clones of Myzus persicae and two clones of the potato aphid, Macrosiphum euphorbiae.The clones of M. persicae showed very different levels of aggressiveness on lettuce: two had a high level of reproduction, two had an intermediate level and two were poorly adapted to lettuce as a host. Differences between lettuce lines in aphid reproduction increased with increasing aggressiveness of the aphid clone, which means that aggressive clones are most effective for selection purposes. No evidence was found for clone-specific plant genotype reactions, meaning that lines resistant to one clone will also be resistant to other clones of M. persicae, allthough not neccessarily at the same level. The lettuce lines selected for partial resistance to the aggressive clone WMp1 were completely or almost completely resistant to less aggresive clones.No differences in level of reproduction were found between the two clones of M. euphorbiae and no relation was observed between resistance to M. persicae and M. euphorbiae, indicating the species-specific character of resistance to leaf aphids in lettuce.     

The response to saprol (systemic fungicide) in lettuce species and cultivars and its inheritance

Summary The response of Lactuca sativa, L. serriola and L. saligna to saprol was studied. Spraying the whole plant or part of a leaf caused wilting of susceptible plants. A similar effect was observed when leaves were soaked in a solution containing saprol.Inheritance studies with L. sativa and L. serriola showed that resistance to the wilting reaction caused by saprol is controlled by a single recessive gene.Contribution no. 198-E, 1978 series.     

Heterosis in crisphead lettuce (Lactuca sativa L.) hybrids

Summary Six of 16 F₁ hybrids from diverse crosses of crisphead lettuce with butterhead, cos or other crispheads significantly outyielded their respective better parent. Four were crosses between crisphead and cos parents, one was a butterhead × crisphead cross and one, a winter crisphead × winter crisphead cross. This latter hybrid plus one other also proved heterotic in a subsequent trial of ten crosses involving five winter crisphead parents, with yield excesses over better parent of 16 and 19%. It appeared significant that the heterotic crosses were given by parents of dissimilar origin. Trials of F₂ segregants from one of these heterotic hybrids enabled predictions to be made for 10.2 to 24.1% yield increases over better parent by the retention and clonal multiplication of the 10% highest yielding selections. Heterosis demonstrated in these trials may constitute the first reported cases in lettuce.     

An association between resistance to root aphid (Pemphigus bursarius L.) and downy mildew (Bremia lactucae Regel) in lettuce

Summary Many lettuce cultivars (Lactuca sativa L.) with high resistance to lettuce root aphid (Pemphigus bursarius L.) also carried the gene Dm-6 for specific resistance to downy mildew (Bremia lactucae Regel). This suggests the possibility of linkage between this gene and root aphid resistance. The origin of this association is discussed.     

Inheritance of the resistant reaction of the lettuce cultivar `Grand Rapids' to the southern root-knot nematode Meloidogyne incognita (Kofoid & White) Chitwood

Resistance to the southern root-knot nematode Meloidogyne incognita Chitwood would be an important attribute of lettuce Lactuca sativa L. cultivars adapted to both protected and field cultivation in tropical regions. `Grand Rapids' and a few other cultivars are reported to be resistant to this nematode. In this paper, we studied the inheritance of the resistant reaction of `Grand Rapids' (P₂) in a cross with a standard nematode-susceptible cultivar Regma-71 (P₁). F₁(Regina-71 × Grand Rapids) and F₂ seed were obtained, and inoculated along with the parental cultivars with different races of M. incognita to evaluate nematode resistance. Broad sense heritability estimates for the number of galls and of egg masses per root system, gall size and gall index were generally in the order of 0.5 or higher. Class distributions of these variables over generations P₁, P₂, F₁ and F₂ were in agreement with simulated theoretical distributions based on monogenic inheritance models. F₃ families were obtained from randomly sampled F₂ plants and tested for reaction to the nematode. The frequency ratio of homozygous resistant, segregating and homozygous susceptible F₃ families did not differ from the 1:2:1 ratio expected from monogenic inheritance. M. incognita resistance appears to be under control of a single gene locus. The Grand Rapids allele (for which the symbol Me is proposed) is responsible for the resistant reaction, and shows high (though incomplete) penetrance, variable expressivity and predominantly additive gene action. This revised version was published online in July 2006 with corrections to the Cover Date.     

Resistance of lettuce (Lactuca) to the leaf aphid Nasonovia ribis nigri. 2. Inheritance of the resistance

Summary The inheritance of resistance to Nasonovia ribis nigri in L. sativa was investigated. Parents and F₁ and F₂ populations from crosses between the susceptible cultivar Ravel and two resistant breeding lines were tested. In both breeding lines one dominant gene appeared responsible for resistance.     

Research on the inheritance of fasciation in lettuce (Lactuca sativa L.)

Summary Inheritance of fasciation was investigated in reciprocal crosses between two lettuce varieties: Suzan (non-fasciated) and Noran (fasciated). In glasshouses parental plants, F₁, F₂, and F₃ populations were assessed for fasciation according to a scale from 0 (non-fasciated)-9 (extremely fasciated).The observed environmental variation for fasciation of the homozygous parents was very low or absent, but the environmental variation for the F₁ populations was large. On the basis of this large variation of the heterozygous F₁ plants we supposed that such a variation also occurred in F₂ and F₃ populations.Considerable mortality was observed in all generations, which supposedly occurred at random in parents and F₁ but was non-random in the segregating F₂ and F₃ populations. So, means and variances of F₂ populations will be biased and therefore they were not used for genetical analyses of fasciation.From F₁ and F₃ populations and from some F₂ populations it appeared that no differences occurred between reciprocals. The differences for fasciation between reciprocal F₂ populations in the 1977 A experiment may result from non-random plant survival. A regression of F₃-means on the values for fasciation of parental F₂ plants, adjusted for their inbreeding, resulted in a realized h²-narrow of 0.4. This indicates that in the surviving plants of the F₂ populations still additive genetic variation was present to select successfully for non-fasciated plants.